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A new species of Piaractus (Characiformes: Serrasalmidae) from the Orinoco

Basin with a redescription of Piaractus brachypomus
Article in Journal of Fish Biology · April 2019

DOI: 10.1111/jfb.13990
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A new species of Piaractus (Characiformes: Serrasalmidae) from the Orinoco Basin
with a redescription of Piaractus brachypomus
M. D. ESCOBAR1,2 | R. P. OTA3 | A. MACHADO-ALLISON4,5 | | I. P. FARIAS1 | T.
HRBEK1*
Accepted Article
1
Universidade Federal do Amazonas, Departamento de Genética, Instituto de Ciências
Biológicas, Laboratório de Evolução e Genética Animal, Av. General Rodrigo Octávio, 6200,
Coroado I., Manaus, Amazonas, Brasil.

2
Universidade Federal do Amazonas, Programa de Pós-Graduação em Ciências Pesqueiras
nos Trópicos, Av. General Rodrigo Octávio, 6200, Coroado I., Manaus, Amazonas, Brasil.
3
Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Biologia de
Água Doce e Pesca Interior, Av. André Araújo, 2.936 Petrópolis, Manaus, Amazonas, Brasil.
4
Universidad Central de Venezuela, Instituto de Zoología Tropical, Laboratório de
Sistemática de Peces, Ciudad Universitaria de Caracas, Los Chaguaramos, Caracas,
Venezuela
5
Wesleyan University, College of the Environment, 45 Wyllys Ave, Middletown,
Connecticut, USA
Correspondence
M. D. Escobar, Universidade Federal do Amazonas, Departamento de Genética, Instituto de
Ciências Biológicas, Laboratório de Evolução e Genética Animal, Av. General Rodrigo
Octávio, 6200, Coroado I. Cep: 69080-90, Manaus, Amazonas, Brasil.
E-mail: ptigrinum@yahoo.com
This article has been accepted for publication in the Journal of Fish Biology and
undergone full peer review but has not been through the copyediting, typesetting,
pagination and proofreading process, which may lead to differences between this
version and the Version of Record. Please cite this article as doi: 10.1111/jfb.13990
This article is protected by copyright. All rights reserved.

Funding information
This study was supported by grants from the Conselho Nacional de Desenvolvimento
Cientifico e Tecnológico (CNPq 490682/2010–3 and 482662/2013–1 to T.H.; CNPq/PPG7
5570090/2005–9 and CNPq/CT-Amazon 575603/2008–9 to I.P.F.), Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES 53923790287) PhD fellowship to

Accepted Article
M.D.E.L. and CAPES 12002011001P7 to R.P.O. and CNPq research fellowship to I.P.F.
(306804/2013-3) and T.H. (303646/2010-1).
ABSTRACT
Piaractus orinoquensis, a new species of serrasalmid fish, is described from the Orinoco
River basin. The new species differs from congeners by having a slenderer body, relatively
smaller head and snout, more compressed mid-body, fewer scales above and below the lateral
line and diagnostic molecular characters in the coI mitochondrial gene region. We also
provide a re-description of Piaractus brachypomus, restricting its geographic distribution to
the Amazon River basin. Both species are economically important in their respective basins
and need to be independently managed as distinct species.
KEYWORDS
Amazon Basin, meristics, mitochondrial DNA, morphology, serrasalmid
1 | INTRODUCTION

Piaractus brachypomus was originally described by Cuvier (1818) as Myletes brachypomus
from the holotype (MNHN A-8627), collected in Brazil from an unspecified location.
Following a convoluted taxonomic history, Eigenmann (1903) erected the genus Piaractus
with P. brachypomus as the type species, then Norman (1929) transferred P. brachypomus to
the genus Colossoma Eigenmann & Kennedy 1903. Schultz (1944) followed Norman’ s
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classification and reported the geographic distribution of the species as including the
Orinoco, Amazon and La Plata (Paraná-Paraguay) Basins. Schultz (1944) probably included
Piaractus mesopotamicus (Holmberg 1887), which was then known as Colossoma mitrei
(Berg 1895), within his treatment of P. brachypomus. Subsequently, Machado-Allison
(1982) in his study of Venezuelan serrasalmids, revalidated the genus Piaractus Eigenmann
1903 and its sole species P. brachypomus, limiting its distribution to the Orinoco and
Amazon River basins. This proposal was based on specimens from the Orinoco River basin
and a review of occurrences reported in literature (Humboldt & Valenciennes, 1821) that
identified and registered the occurrence of the species in the Orinoco River (Venezuela) and
the upper Marañon River (Peruvian Amazon) and Goulding (1979, 1980) who registered this
species in the Madeira River (Brazilian Amazon).
Géry (1985) highlighted the extensive range of intraspecific phenotypic variability,
raised the possibility of cryptic species and discussed the taxonomy and distribution of P.
brachypomus and P. mesopotamicus. However, in spite of Géry’ s (1985) observations, all
subsequent taxonomic and molecular studies (Jégu, 2003; Ortí et al., 2008; Oliveira et al.,
2011) and biological inventories (Taphorn, 2003; Machado-Allison et al., 2005; Watkins et
al., 2005; Galvis et al., 2006; Maldonado-Ocampo et al., 2006; Ortega et al., 2006; Santos et
al., 2006; Agudelo et al., 2011; Lasso & Sánchez-Duarte, 2011) have followed the taxonomy
and distribution of P. brachypomus proposed by Machado-Allison (1982). Escobar et al.

(2015) were the first to recognize that P. brachypomus of the Orinoco and the Amazon
Basins actually represent distinct and independent evolutionarily lineages and therefore
different species.
A relatively large number of species are thought to be shared between the Amazon and
Orinoco River basins both due to shared geological history and their current connection via
the Casiquiare Canal (Lundberg et al., 1998). However, Winemiller & Willis (2011) pointed
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out that the Casiquiare Canal acts more often as an ecological filter of aquatic species than a
dispersal route and thus, it is likely that the number of species shared between the Orinoco
and Amazon Basins may be smaller than previously thought. Biogeographical studies also
indicate little similarity between the two basins (Hubert & Renno, 2006; Dagosta & Pinna,
2017), which further supports a pattern of greater faunal dissimilarity than traditionally
thought.
Traditionally, species descriptions are based primarily on morphological characters,
with secondary support provided by ecological and life-history characters (Winston, 1999).
Given the increasingly easy accessibility of molecular techniques to taxonomists, an
increasing number of species descriptions are now including molecular data (Benine et al.,
2009; Melo et al., 2016; Andrade et al., 2017). In this study we use multiple lines of evidence
(sensu Padial et al., 2010) to describe a new species of the genus Piaractus from the Orinoco
River basin. Species delimitation approaches that unite multiple lines of evidence (Dayrat,
2005) can provide robust taxonomic hypotheses, especially in morphologically conserved
groups; i.e., taxonomic groups harbouring cryptic or pseudocryptic taxa. Conceptually this
approach views species as evolving lineages, general lineage species concept sensu de
Queiroz (2007), that exhibit the properties of independently evolving lineages, such as
reciprocal monophyly, mutually exclusive mating systems and diagnosability. In addition to

the description of the new species from the Orinoco Basin, we also provide a re-description
of P. brachypomus and we delimit the geographic distribution of both species.
2 | MATERIAL AND METHODS

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2.1 | Fish samples
For molecular analyses, specimens were collected between 2010 and 2013 from five
localities in the Orinoco Basin (Guaviare River in San José del Guaviare; Meta River in
Puerto López, Orinoco River in Puerto Carreño; Apure River in San Fernando de Apure and
Orinoco River in San Felix) and from 11 localities in the Amazon Basin (Solimões–Amazon
River in Leticia, Tefé, Janauacá and Santarém; Japurá River in La Pedrera and Japurá;
Tapajós River in Itaituba; Madeira River in Guarará-Mirim, Humaitá and Borba; Parimé
River tributary of the Branco River in Roraima). Representative specimens were deposited at
the ichthyology reference collection of the Guyana Hydrobiological Research Station, La
Salle Foundation (CIR-EDIHG), San Félix, Venezuela, under the lot numbers CIR-EDIHG
8195 and 8196 and at the National Institute of Amazonian Research (INPA), Manaus, Brazil
under the lot number 39188. Additional examined material came from the Philadelphia
Academy of Natural Sciences (ANSP), Philadelphia, USA and Museum of Biology of the
Central University of Venezuela (MBUCV-V), Caracas, Venezuela.
The distribution map for both species was generated using the program QGIS 2.8
(QGIS Development Team, 2018).
2.2 | Molecular methods

Molecular data are from Escobar et al. (2015) and consist of 145 mitochondrial DNA control
region (d-loop) sequences from 60 individuals from the Orinoco Basin and 85 individuals
from the Amazon Basin (GenBank KP705146–KP705254) and 33 cytochrome c oxidase
subunit I (coI) sequences from 19 individuals from the Orinoco Basin and 14 individuals
from the Amazon Basin (Genbank KP723341–KP723360, MK395376– MK395388).
The sequences were edited in the software BioEdit (Hall, 1999) and aligned in Clustal
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W (Thompson et al., 1997). We used population aggregation analysis (PAA) performed at the
level of an individual (Davis & Nixon, 1992; Rach et al., 2008) as implemented in the SPIDER
package in R (Brown et al., 2012; www.r-project.org) to identify diagnostic sites in the coI
gene. The control region sequences were visualized as a haplotype network generated in the
software Haploviewer (Salzburger et al., 2011) and a maximum-likelihood phylogeny
inferred in the software RAxML (Stamatakis, 2014) using the GTRCAT model of molecular
evolution.
We also inferred a maximum-likelihood phylogeny from the coI sequence alignment in
the software RAxML (Stamatakis, 2014) using the GTRCAT model of molecular evolution,
with phylogenetic support being assessed via 1000 bootstrap replicates. As immediate
outgroups we used Colossoma macropomum (Cuvier 1816) (MG751942) and Mylossoma
albiscopum (Cope 1872) (MG752455) and we included P. mesopotamicus as additional
ingroup.
Finally, we used mPTP, the multi-rate Poisson tree process (Kapli et al., 2017),
coalescent single-locus species-discovery method that has been designed specifically to
differentiate between population structure and species-level divergence to test for the
presence of multiple species of Piaractus in the Orinoco and Amazon Basins. We first
reduced the dataset to unique haplotypes using the function hapCollapse
(www.github.com/legalLab/protocols-scripts) to further minimise interpreting population-

level structuring as species-level differentiation (Talavera et al., 2013) and then inferred a
maximum-likelihood phylogeny from the coI sequence alignment in the software RAxML
(Stamatakis, 2014) using the GTRCAT model of molecular evolution. The resulting
phylogeny was analysed in the software mptp 0.2.3 (Kapli et al., 2017).
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2.3 | Morphological methods
Measurements were taken point-to-point with a Mitutoyo Absolute digital callipers (model
500-172-20B, CD-8” CX-B; www.mitutoyo.co.uk) to the nearest 0.1 mm and counts under a
Zeiss stereomicroscope (www.zeiss.com), both on the left side whenever possible, following
standard methods proposed in Machado-Allison (1982). All distances were calculated in mm
and transformed to percentages; the body measurements are expressed as a proportion of
standard length (LS), while head measurements are expressed as a proportion of head length
(LH). Relative distances did not deviate from normal expectations. We therefore applied a t-
test in addition to the Mann-Whitney U-test to test for differences between fishes of the two
basins. These analyses were performed in the STATS package in R.
Vertebral and supraneural elements were counted in 17 skeletonised specimens [eight
of P. brachypomus: INPA 46334 (1 skeleton), 46335 (1 skeleton), 50841 (4 skeletons), 57799
(2 skeletons)] and nine of the new species [INPA 57798 (2 skeletons) and ICN-MHN 24051
(4 skeletons)] and three x-rayed specimens (one P. brachypomus, ANSP 165052 and two of
the new species, ANSP 163570 and 189567), designated as skeleton and x-rayed in the
species accounts. Dried skeletons were cleaned with the aid of dermestid beetle larvae and X-
ray radiographs were taken in the Faxitron specimen radiography system LX-60
(www.faxitron.com). Vertebral counts include the Weberian apparatus as four elements and

the fused PU1+U1 as a single element. Osteological terminology follows Weitzman (1962)
and Mattox et al. (2014).
In the descriptions, counts are followed by their frequency in parentheses and an
asterisk indicates values for the holotype. Ranges of the new species are followed by the
values of the holotype in parentheses. In the list of analysed material, total number of
specimens of each lot is given first, followed by those skeletonised or X-rayed. Institutional
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abbreviations are: IAvH, Instituto de Investigación de Recursos Biológicos Alexander von
Humboldt, Villa de Leyva, Boyacá, Colombia; ICN-MHN, Instituto de Ciencias Naturales,
Museo de História Natural, Universidad Nacional de Colombia; UFRO, Universidade Federal
de Rondônia, Laboratório de Ictiologia e Pesca, Porto Velho, Brazil; CIR-EDIHG, INPA and
MBUCV-V, aforementioned.
For geometric morphometric analyses, 14 landmarks (Supporting Information Figure
S1) were collected from 33 specimens, 12 from the Amazon and 22 from the Orinoco Basins
using the software tpsDIG 2.17 (Rohlf, 2013). We then carried out a generalised Procrustes
transformation followed by visualisation on a thin-plate-spline grid of the transformation of
the mean shape in each basin to the mean shape in the other basin. The partial warp scores of
the Procrustes residual of the Orinoco and Amazon specimens were compared using
Hotelling’ s T2. All transformations and analyses were carried out in the shapes package in R
(Dryden, 2018).
3 | RESULTS
Results of all analyses consistently indicate that Piaractus from the Orinoco and Amazon
Basins are different species. Piaractus from the Amazon Basin has a significantly deeper
body and longer maxilla than Piaractus from the Orinoco Basin (Figure 1 and Table 1). The

two groups are diagnosable by non-overlapping counts of lateral-line scales, scale rows
below and above the lateral line (Table 1) and by a series of molecular apomorphies in the
mitochondrial coI gene (Table 2). The mPTP (1-methyl-4-phenyl-1,2,3,6-
tetrahydropyridine) coalescent-based single locus species discovery method delimits the
fishes from the Orinoco and Amazon Basins as two different species. Finally, Orinoco and
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Amazon populations form distinct reproductive groups that are not connected by geneflow
and that diverged from each other approximately 2.54 M years ago (Escobar et al., 2015). All
these lines of evidence lead us to conclude that Piaractus from the Orinoco and Amazon
Basins are different species.
3.1 | Piaractus orinoquensis sp. nov.
Figures 1-3 and Table 1-3
Colossoma brachypomus Schultz 1944: p. 255 (río Caripe, cerca Caripito, Venezuela), Mago-
Leccia, 1970: p. 70 (citation); Colossoma brachypomum (Cuvier 1818) Novoa, 1982: p. 268
(biology); Piaractus brachypomus (Cuvier 1818) Machado-Allison, 1982: p. 47 (detailed
description, ontogenetic development, distribution), Machado-Allison & Fink, 1995: p. 84
(description, distribution), Taphorn et al., 1997: p. 74 (citation); Novoa, 2002: p. 143
(description, habitat, life history, migration, fishing aspects); Jégu, 2003: p. 187 (citation,
synonymy, distribution); Taphorn, 2003: p. 66, 220-222 (identification key, synonyms,
description, distribution, live history), Lasso et al., 2004: p. 156 (citation), Machado-Allison
et al., 2005: p. 197 (citation), Maldonado-Ocampo et al., 2008: p. 174 (citation), Usma et al.,
2009: p. 105 (citation, migration notes), Agudelo et al., 2011: p. 236 (citation, figure,
diagnosis, geographic distribution, life history, fisheries information, conservation status),

Lasso & Sanchez-Duarte, 2011: p. 131 (diagnosis, life history, fisheries information,
distribution, figures 96a,b), Machado-Allison et al., 2013: p. 294 (citation, figure 3c).
3.1.1 | Holotype
ANSP 163570, 150.2 mm LS, X-rayed, Laguna La raya, caño Falcón, nearby Camaguan,
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Guárico, 8° 07’ N, 67° 37’ W, B. Chernoff & J. Lundberg, 22 January 1988 (Figure 2).
3.1.2 | Paratypes
All from Orinoco River Basin.
Venezuela, Apure River, Apure: MBUCV-V 607, 2, 116.0-121.0 mm LS, Laguna El
Pavoncito, San Fernando de Apure, 7° 51’ 50” N, 67° 35’ 40” W, 16 January 1963, J. A.
Fernández & J. A. Luengo; MBUCV-V 634, 1, 139.0 mm LS, Caño Los Zorros and beach
near Los Bosquerones, San Fernando de Apure, c. 7° 55’ N, 67° 37’ W, 17 January 1963, F.
Mago Leccia & J. A. Luengo; MBUCV-V 14002, 1, 153.0 mm LS, Apure River close to San
Fernando de Apure, 7° 54’ N, 67° 29’ W, 15 May 1978, J. A. Luengo & L. Aguana;
MBUCV-V 14003, 2, 40.0–45.0 mm LS, Las Maporas, flooded zone near San Fernando de
Apure, 7° 49’ 57” N, 67° 24’ 24” W, 26 January 1978, A. Ríos; MBUCV-V 14004, 3, 47.3–
59.0 mm LS, Caramacate nearby San Fernando de Apure, 7° 51’ N, 67° 27’ W, 27 August
1978, A. Ríos; MBUCV-V 14005, 6, 25.4-45.0 mm LS, La Guanota, nearby San Fernando de
Apure, 7° 53’ N, 67° 26’ W, 17 July 1979, L. Aguana, R. Gill & A. Machado; MBUCV-V
14006, 6, 29.0–36.8 mm LS, Manglarote, vía Arichuna San Fernando de Apure, 7° 53’ N, 67°
25’ W, 18 July 1979, A. Machado-Allison, R. Gill & L. Aguana; MBUCV-V 23586, 10,

118.0–185.0 mm LS, lagunita del río Apurito, 7° 55’ 56” N, 68° 28’ 05” W, 29 October 1976,
T. Zaret, H. Lopez & L. Aguana.
Guárico River, Guárico: MBUCV-V 305, 116.6 mm LS, San Jose River, tributary of Guárico
River, 8° 14’ 50” N, 66° 59’ 54” W, 25 May 1960, M. V. Ramírez,; MBUCV-V 6641, 1,
121.0 mm LS, río Aguaro nearby Santa Rita, vía Las Mercedes-Cabruta, 8° 08’ 47” N, 66°
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14’ 01” W, 22 November 1968, J. Moscó, H. Mendoza, A. Cortéz & A. Machado-Allison.
Orinoco River, Bolívar: ANSP 166684, 2, X-rayed, 116.6–142.6 mm LS, lagoon tributary of
Orinoco River, Almacen, 8° 06’ N, 63° 45’ 50” W, 21 January 1987, M. Rodriguez; INPA
58179, 2, 118.7–130.4 mm LS, same data as ANSP 166684; ANSP 166685, 1, X-rayed, 89.8
mm LS, Orinoco River, Cabruta, 7° 38’ 55” N, 66° 13’ 40” W, 12 January 1987, M.
Rodriguez; CRI-EDIHG 8195, Genbank KP723347, 2, 271.0–245.0 mm LS, Isla de
Providencia, near Isla Fajardo, confluence of the Caroni and Orinoco Rivers, Guayana, 8° 20’
03” N, 62° 42’ 16” W, November 2013, M. D. Escobar, V. Verona and N. González; CIR-
EDIHG 8196, Genbank KP723348, 299.0 mm LS, same data as CRI-EDIHG 8195; MBUCV-
V 1554, 2, 57.2-61.3 mm LS, Laguna del Medio, Ciudad Bolívar, 8° 08’ 49” N, 63° 31’ 44”
W, 7 September 1963, F. Mago Leccia; MBUCV 5380, 1, 131.5 mm LS, Caño Cocuiza,
puente Tabirito, Caicara del Orinoco, 7° 39’ N, 66° 05’ W, 27 February 1968, F. Mago
Leccia, J. Moscó & A. Machado-Allison.
Monagas: MBUCV-V 1676, 2, 96.0–103.0 mm LS, Laguna El Guatero, Barrancas, 8° 41’ 02”
N, 62° 12’ 08” W, 11 November 1963, F. Mago Leccia & E. Miranda.

Delta Amacuro: MBUCV-V 19186, 6, 84.0–109.0 mm LS, Caño Milla 49 km from sea, Delta
of Orinoco River, 8° 39’ N, 61° 08’ W, 20 November 1979, J. Baskin & D. Stewart.
Portuguesa River, Guárico: ANSP 189567, 1, X-rayed, 110.9 mm LS, caño Falcón, north of
San Fernando de Apure, 8° 07’ N, 67° 37’ W, 26 November 1976, J. Lundberg et al.;
MBUCV-V 5629, 1, 103.0 mm LS, Caño Corozo Pando, on road Calabozo-San Fernando de
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Apure, 8° 14’ N, 67° 42’ W, 15 October 1968, J. Moscó, O. Travieso & H. Mendoza;
MBUCV-V 6570, 1, 218.0 mm LS, flooded lagoon nearby caño Falcón, north of Camaguán,
8° 07’ N, 67° 37’ W, 16 July 1971, O. Silva, A. Machado-Allison & L. Aguana; MBUCV-V
6613, 3, 140.0–166.0 mm LS, Laguna Los Laureles, between Camaguán and caño Falcón, 8°
06’ N, 67° 37’ W, 20 March 1972, A. Machado-Allison & L. Aguana; MBUCV-V 6661, 3,
185.0-188.0 mm LS, same data as MBUCV-V 6613, 7 May 1972; MBUCV-V 7057, 3, 86.0–
127.0 mm LS, same data as MBUCV-V 6613, 15 November 1972, L. Aguana & O. Silva;
MBUCV-V 7146, 3, 117.0–134.5, mm LS, same data as MBUCV-V 6613, 15 November
1972; MBUCV-V 7087, 1, 102.5 mm LS, same data as MBUCV-V 6613, 15 November 1972;
MBUCV-V 23851, 9, 67.5–130.0 mm LS, Préstamo Carretera close to Caño Falcón, 8° 05’ N,
67° 37’ W, 23 January 1979, J. Luengo & L. Aguana.
Colombia, Meta River, Puerto López: INPA 57798, Genbank MK395376-80, 5 skel, 82.0–
145.5 mm LS, Meta River, Puerto López, Meta, 4° 08’ 11” N, 72° 55’ 46” W, January 2018,
M. D. Escobar L; ICN-MHN 24051, Genbank MK395381-4, 4 skel, 85.0-143.6 mm LS, same
data as INPA 57798; unvouchered, Genbank MK395385-6, same data as INPA 57798.
3.1.3 | Morphological diagnosis

Piaractus orinoquensis sp. nov., differs from P. brachypomus and P. mesopotamicus by
possessing fewer rows of scales above (23–30 v. 31–37 and 50–60, respectively) and below
the lateral line (22–32 v. 33–37 and 49–56, respectively). Additionally, the minimum number
of lateral-line scales helps to distinguish the new species from its congeners (72–89, mode 82
v. 84–100, mode 87 in P. brachypomus and 107–128, mode 117 in P. mesopotamicus).
Piaractus orinoquensis sp. nov. has a significantly shallower body when compared with P.
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brachypomus (Hotelling T2 = 14.95836, P < 0.001). Specifically, Piaractus orinoquensis sp.
nov. is significantly shallower at mid-body and the anterior edge of the caudal peduncle and
the distance between snout and the posteriormost supraoccipital is significantly shorter (Table
1 and Figure 1). The maxilla is also significantly shorter than in P. brachypomus (Table 1 and
Figure1).
3.1.4 | Molecular diagnosis
Piaractus orinoquensis sp. nov. is diagnosable from P. brachypomus by the presence of 34
diagnostic molecular character states in d-loop position and 19 diagnostic molecular
character states in the coI gene (Table 2).
3.1.5 | Description
Morphometric data are summarised in Table 3. Body compressed, overall aspect of body
ovoid, body deepest at vertical through pelvic-fin origin. Pre-dorsal and postdorsal origin c.
equal in length. Dorsal profile of head convex from mouth to vertical through anterior
portion of eye, concave from latter point to base of supraoccipital process and convex from
that point to dorsal-fin origin. Dorsal body profile steep from snout to dorsal-fin origin and

with slight posteroventral slope after this point. Dorsal-fin base convex. Body profile almost
straight from dorsal-fin insertion to adipose-fin origin. Adipose-fin base convex. Head
robust, with posterior margin surrounded by broad opercle membrane, covering up to three
scales horizontally at point of greatest head length. Ventral keel with 16 to 38 (25) simple
pre-pelvic spines followed by 15 to 28 (20) simple postpelvic spines and three to eight (6)
paired spines around anus; totalling 40 to 66 (51) spines, composing ventral keel, always
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anterior of anus. Caudal peduncle relatively short; upper and lower profile of caudal
peduncle slightly concave.
Snout blunt and rounded. Mouth terminal to slightly upturned, with molariform teeth.
Two rows of premaxillary teeth with five molariform teeth in labial row and two in lingual
row. Teeth in labial row gradually decreasing in size posteriorly. Labial and lingual
premaxillary teeth rows interspaced by gap. Dentary with six to seven (6) teeth decreasing in
size posteriorly; pair of symphyseal dentary teeth always present behind main series of teeth.
Maxillary with none to three small teeth (0).
Scales cycloid, irregular in sized but relatively large for serrasalmid species. Lateral
line complete with 72 to 89 (80) perforated scales; first six or eight scales larger than the
remaining. Scales rows above lateral line 23 to 30 (29). Scales rows below lateral line 22 to
32 (25). Dorsal-fin rays ii to v (iv), 13 to 15 (14); not reaching adipose-fin origin when
adpressed. Dorsal-fin origin at mid-body or slightly posterior of vertical through pelvic-fin
origin; dorsal-fin margin truncate. Adipose fin with distal margin rounded to subrectangular
with no rays. Pectoral-fin rays i, 15 to 18 (16), with anterior rays longer than others. Pelvic-
fin rays i, 7 (7), anterior rays slightly more elongate than others, not reaching anal-fin origin.
Anal-fin rays ii to iv (iv), 22 to 25 (23), with anterior rays longer than others, falcate distal
margin. Caudal fin forked, with lobes of similar size. First branchial arch with short gill

rakers, 16 to 19 (16) gill rakers on upper limb; 15 to 20 (17) gill rakers on lower limb; one at
cartilage between upper and lower limbs.
Premaxilla strongly attached to mesethmoid. Ascending premaxillary process elongate
with similar width from base to distal tip, distal tip rounded and moderately inclined in
relation to lateral premaxillary process. Lateral premaxillary process triangular, with
dorsolateral process with concavity for maxillary insertion. One large crypt of replacement
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teeth on premaxilla. Ascending arm of maxilla narrow and pointed, descending arm of
maxilla expanded and large. Dentary rectangular and slightly arched with one large bony
replacement tooth crypt. Symphyseal dentary teeth well-developed and surpassing dorsal
edge of first tooth from main row. Antorbital narrow almost triangular, with base wider.
Supraorbital narrow, with regular margins, not contacting infraorbital 5. Five well-developed
infraorbitals forming semi-circle and leaving narrow naked area on cheeks. Infraorbitals 3
and 4 with branched sensory canal and 5 with Y-shaped canal. Broad orbital region. Total
vertebrae 33 to 35 (38); abdominal vertebrae 14 or 15 (15); caudal vertebrae 18 to 20 (18).
One vertebra between verticals through last dorsal-fin pterygiophore and first anal-fin
pterygiophore. Supraneurals 5 or 6 (6).
3.1.6 | Colour in alcohol
Overall body colouration brown to yellow, dorsal region of body darker. Flanks with
inconspicuous rounded or vertically elongate dark blotches, most concentrated above the
lateral line and just below it (Supporting Information Figure S2). Opercle with a
conspicuous dark blotch immediately below mid-region of the bone. Eye with broad dark
transversal bar crossing pupil (not discernible in specimens retained for a long period in
formalin). Dorsal fin hyaline. Adipose and caudal fins c. hyaline and darker towards fin

margins, with chromatophores along distal margin forming narrow dark stripe. High
concentration of dark pigments on middle caudal-fin rays. Pectoral, pelvic and anal fins
dark. Pectoral and pelvic fins with pigments most concentrated on branched rays and darker
towards fins tip. Anal fin with dark chromatophores along interradial membranes. Early-
stage juveniles (below 60 mm LS) with mid-lateral ocellus (6.7 to 10.1% of LH) and with
dorsal, adipose, caudal and anal fins hyaline with chromatophores along distal margin
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forming narrow dark stripe. Pectoral and pelvic fins completely hyaline.
3.1.7 | Colour in life
Body with intense silver colour on dorsum, upper part of the head and sides of body.
Inferior portion of head including preopercle, sub-opercle, inferior portion of infra-orbitals,
opercle and gular, abdominal and supracleithrum region yellow to orange hues. Opercle
with a dark conspicuous blotch on lower half, less conspicuous in exemplars longer than 300
mm LS (Supporting Information Figure S3). Dorsal, adipose and caudal fins hyaline, with
dark chromatophores concentrated on distal portion of fins, forming thin dark margin.
Pectoral, anal and posterior portion of pelvic fins orange. Anal fin with a dark distal margin.
Caudal fin with wide distal black band. Juveniles and young adults with strong orange tones
on inferior part of the head, ventral and inferior region of flanks, making them quite similar
in colouration to the red-belly piranhas (Pygocentrus spp.). This tone intensity decreases in
large adult individuals. Early-stage juveniles with mid-lateral ocellus with orange
colouration described previously in colour in alcohol.
3.1.8 | Distribution

Piaractus orinoquensis sp. nov. is normally present in white, semi-clear and occasionally in
the interface of black-white water tributaries of the Orinoco River. In the western region, it
can be found in the Guaviare, lower Inírida, Vichada, Tuparro, Tomo, Bita, Meta (including
tributaries such as the Manacacias, Casanare, Cravo Norte) and Arauca Rivers. North of the
Orinoco River it can be found in the Arichuna, Capanaparo, Cinaruco, Guariquito, Apure
(including tributaries such as the Uribante, Sarare, Caparo, Suripa, Ticoporo, Canaguá,
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Pagüey, Masparro, Santo Domingo, Portuguesa and Guárico), Manapire, Zuata, Pao and
Caris Rivers. To the south of the Orinoco in the Atabapo, Sipapo, Parguaza, Suapure,
Cuchivero, lower Caura, lower Aro and lower Caroní Rivers and in the delta of the Orinoco
River itself (Figure 3).
3.1.9 | Etymology
The species name, orinoquensis, refers to the geographic area where the new species is
known, from Orinoco River basin. A noun in apposition.
3.2 | Piaractus brachypomus (Cuvier 1818)
Figures 2–4 and Tables 1, 2, 4
Myletes brachypomus Cuvier, 1818: p. 452 (original description, type locality: Brazil,
holotype: MNHN A.8627), Cuvier & Valenciennes, 1849: p. 199 (redescription of the
holotype); Myletes paco Humboldt & Valenciennes, 1821: p. 175 (original description, type
locality: Guaporé River, Amazon River basin, holotype: unknown); Myletes bidens Agassiz
in Spix & Agassiz 1829: p. 75 (original description, type locality: rivers of equatorial Brazil,
holotype: unknown), Géry, 1985: p. 98 (designation of lectotype: MNHN 789); Salmo pacu

Kner 1860: p. 21 (name in text); Piaractus brachypomus Eigenmann, 1903: p. 148 (new
combination), Machado-Allison, 1982: p. 47 (detailed description, ontogenetic development,
distribution), Ortega et al., 2012: p. 37 (occurrence in Amazon River in Peru), Galvis et al.,
2006: p. 199 (occurrence in Amazonas River in Colombia), Santos et al., 2006: p. 51
(diagnosis, biology, fisheries information), Maldonado-Ocampo et al., 2008: p. 174 (citation),
Usma et al., 2009: p. 105 (citation, migration notes), Agudelo et al., 2011 (citation, figure,
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diagnosis, geographic distribution, life history, fisheries information, conservation status),
Barriga 2011: p. 108 (occurrence in Ecuador), Zeinad & Prado 2012: p. 162 (description,
diagnosis, life history, distribution, fisheries information), Ota et al., 2013: pp. 28-29 (brief
description, diagnosis, occurrence in Madeira River); Reganina bidens Fowler, 1906: p. 475
(allocated in Regania); Colossoma bidens Eigenmann, 1910: p. 444 (cited); Norman, 1929: p.
810 (identification key, redescription), Goulding, 1979: p. 57 (life history, fisheries
information, migration); Colossoma brachypomus Norman, 1929: p. 809 (synonymy),
Fowler, 1950: p. 388 (synonymy, distribution), Waiteina fowleri Campos, 1946: p. 219
original description, type locality rio Tapajós, Brazil, Holotype: MZUSP (ex DZSASP) 357).
3.2.1 | Material examined: type specimen
MNHN A.8627, holotype of Myletes brachypomus, 545.0 mm LS, dry, Brazil, without exact
location, Cuvier, 1818 (Figure 4).
3.2.2 | Non-type specimens
Brazil, Amazon River basin, Amazonas: INPA 5790, 9, 96.0–172.0 mm LS, ilha de
Marchantaria, rio Solimões, 3° 14’ 57” S, 59° 58’ 22” W, Iranduba, ichthyology team of

INPA, 2 December 1991; INPA 25754, 1, 101.0 mm LS, lago Maratu, 60 km from the
confluence of the rios Negro and Solimões, Manaus, 3° 20’ 51” S, 60° 12’ 34” W, M. C.
Vega, April 2003; INPA 39188, Genbank KP723355-6, 2, 202.0–242.0 mm LS, lago
Janauacá, rio Amazonas, Careiro, ca. 3° 27’ S, 60° 16’ W, M. D. Escobar, 8 February 2013;
INPA 46334, 1 skeleton, 101.3 mm LS, lago do Padre, complex of Catalão lake, Iranduba, 3°
10’ 09” S, 59° 54’ 43” W, S. A. Amadio, 2 July 2014; INPA 46335, 1 skeleton, 155.9 mm LS,
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same data as INPA 46335.
Branco River basin, Roraima: INPA 23435, 1, 141.3 mm LS, rio Branco, Boa Vista, 2° 43’
28” N, 60° 42’ 08” W, V.N. Machado & T. Hrbek, 10 October 2013.
Madeira River basin, Rondônia: INPA 3817, 1, 95.0 mm LS, rio Madeira, Calama, 8° 01’ 25”
S, 62° 52’ 38” W, M. Goulding, 30 September 1980; INPA 10985, 2, 79.0–104.0 mm LS, rio
Guaporé at its confluence with rio Mamoré, Guajará-Mirim, 10° 29’ 51” S, 65° 24’ 57” W,
G. M. Santos, 19 June 1984; INPA 38551, 1, 99.0 mm LS, lago do Bodó, comunidade Bom
Jardim, Porto Velho, 8° 32’ 33” S, 63° 37’ 28” W, L. Costa, F. Vieira, A. Negrão et al., 12
August 2001; INPA 38643, 2, 1c&s, 108.0–110.2 mm LS, lago redondo na comunidade São
Salvador, rio Madeira, Humaitá, 6° 43’ 04” S, 62° 19’ 34” W, L. Costa, F. Vieira & A.
Negrão, 23 Ago 2011; INPA 44003, 1, 224.0 mm LS, rio Madeirinha, Autazes, 3° 30’ 39” S,
59° 03’ 41” W, Normando, F.T. & Nunes, D.M.F., 8 November 2013; INPA 53765, 1, 29.3
mm LS, rio Lajes, Nova Mamoré, 10° 25’ 46” S, 65° 22’ 18” W, Naturae team, 8 May 2012;
INPA 54505, 1, 300.0 mm LS, rio Abunã, Porto Velho, 9° 41’ 38” S, 65° 26’ 42” W, Naturae
team, 11 February 2011; UFRO 6289, 315.0 mm LS, lago Cuniã, Madeira River, Rondônia,
8° 17’ 13” S, 63° 28’ 48” W, ichthyology team of INPA, 10 April 2010.

Negro River basin, Amazonas: INPA 50841, 101.8–114.8 mm LS, 4 skeleton, rio Negro,
Anavilhanas archipelago, Novo Airão, 2° 33’ 31” S, 60° 55’ 24” W, E. G. Ferreira, 12 June
2014.
Tapajós River basin, Mato Grosso: INPA 48524, 1, 583.0 mm LS, rio Teles Pires,
downstream 7 Quedas rapids, Paranaita, 9° 19’ 17” S, 56° 46’ 41” W, W. Ohara, February
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2014.
Tocantins River basin, Pará: INPA 3814, 3, 100.0–106.0 mm LS, rio Tocantins, Icangui, ca.
3° 38’ S, 49° 39’ W, ichthyology team of INPA, 29 June 1980; INPA 3816, 1, 92.0 mm LS,
rio Tocantins, Itupiranga, ca. 5° 03’ S, 49° 19’ W, ichthyology team of INPA, 4 July 1980 ;
INPA 20122, 2, 146.0–191.0 mm LS, rio Tocantins, praia do Coco, Parque Estadual do
Cantão, ca. 9° 14’ S, 49° 59’ W, Cantão project team, 17 November 2000; INPA 20628, 1,
194.1 mm LS, rio Tocantins, Baião, 2° 50’ 17” S, 49° 40’ 54” W, G. M. Santos, 24 May
2000; INPA 20629, 1, 335.8 mm LS, rio Tocantins, Icangui, 3° 17’ 40” S, 49° 38’ 42” W, G.
M. Santos, 25 April 2001.
Colômbia, Amazonas: IAVH 11909, 1, 145.0 mm LS, lago Tarapoto, Pozo de Zoraida, Puerto
Nariño, 3° 47’ 58” S, 70° 27’ 25” W, S. Correa, 2007; IAVH 11570, 1, 154.0 mm LS, same
data as IAVH 11909; IAVH 11571, 1 165.0 mm LS, same data as IAVH 11909; INPA 57799,
Genbank MK395387-8, 2 skeleton, 308.5–310.0 mm LS, Solimões River, Letícia, 4° 09’ 03”
S, 70° 00’ 48” W, M. D. Escobar, January 2018.
Peru, Loreto: ANSP 165052, 1, X-rayed, 86.2 mm LS, Quebrada of Yanayacuríve, W of
Miraflores, río Marañon, 4° 42’ S, 74° 17’ W, P. Fromm et al., 21 August 1989.

3.2.3 | Morphological diagnosis
Piaractus brachypomus differs from P. orinoquensis and P. mesopotamicus by possessing
31–37 rows of scales above lateral line (v. 23–30 and 50–60, respectively) and 33 and 37
rows of scales below lateral line (v. 22–32 and 49–56, respectively). Additionally, an
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intermediate number of lateral line scales helps to distinguish P. brachypomus from its
congeners (84–100, mode 87 v. 72–89, mode 82 in P. orinoquensis and 107–128, mode 117
in P. mesopotamicus). Piaractus brachypomus has a significantly deeper body when
compared with Piaractus orinoquensis (Hotelling T2 = 14.95836, P < 0.001). Specifically, P.
brachypomus is significantly deeper at mid-body and the anterior edge of the caudal peduncle
and the distance between snout and posteriormost supraoccipital is significantly longer (Table
1 and Figure 2). The maxilla is significantly longer than in Piaractus orinoquensis (Table 1;
Figure 2).
3.2.4 | Molecular diagnosis
Piaractus brachypomus is diagnosable by at least 34 molecular character states in d-loop
position and 19 character states in the coI gene (Table 2).
3.2.5 | Description
Morphometric data summarised in Table 4. Body compressed, overall aspect of body ovoid,
body deepest at vertical through pelvic-fin origin. Predorsal and postdorsal origin c. equal in

length. Dorsal profile of head convex from mouth to vertical through anterior portion of eye,
gently concave from latter point to base of supraoccipital process and convex from that point
to dorsal-fin origin. Dorsal body profile steep from snout to dorsal-fin origin and with slight
posteroventral slope after this point. Dorsal-fin base convex. Body profile almost straight
from dorsal-fin insertion to adipose-fin origin. Adipose-fin base convex. Head robust, with
posterior margin surrounded by broad opercular membrane, covering up to three scales

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horizontally at point of greatest head length. Ventral keel with 29-38 simple pre-pelvic spines
followed by 15–37 simple postpelvic spines and three to 11 paired spines around anus;
totalling 48 to 81 spines, composing ventral keel, always anterior of anus. Caudal peduncle
relatively short; upper and lower profile of caudal peduncle slightly concave.
Snout blunt and rounded. Mouth terminal to slightly upturned, with molariform teeth.
Two rows of premaxillary teeth with five molariform teeth in labial row and two in lingual
row. Teeth in labial row gradually decreasing in size posteriorly. Labial and lingual
premaxillary tooth rows interspaced by gap. Dentary with six to seven (7) teeth decreasing in
size posteriorly; pair of symphyseal dentary teeth always present behind main series of teeth.
Maxillary with none to three (2) small teeth.
Scales cycloid, irregular in size though relatively large for serrasalmid species. Lateral
line complete with 84–100 (96) perforated scales; first six or eight scales larger than
remaining. Scale rows above lateral line 31–37(31). Scale rows below lateral line 33–37(33).
Dorsal-fin rays iii,v,13–16(14); not reaching adipose-fin origin when adpressed. Dorsal-fin
origin at mid-body or slightly posterior of vertical through pelvic-fin origin; dorsal-fin margin
truncate. Adipose fin with distal margin rounded to subrectangular. Pectoral-fin rays i,16–
19(16), with anterior rays longer than remaining rays. Pelvic-fin rays i,7(7) with distal margin
convex, anterior rays slightly more elongate than remaining, not reaching anal-fin origin.
Anal-fin rays iii–iv,22–28(23), with anterior rays longer than remaining, falcate distal

margin. Caudal-fin forked, with lobes of similar size. First branchial arch with short gill
rakers, 14–18 gill rakers on upper limb; 15–20 gill rakers on lower limb; one at cartilage
between upper and lower limbs.
Ascending premaxillary process elongate with similar width from base to distal tip,
distal tip rounded and moderately inclined in relation to lateral premaxillary process. Lateral
premaxillary process triangular, with pointed tip. One large crypt of replacement teeth on
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premaxillary. Ascending arm of maxillary narrow and pointed, descending arm of maxillary
expanded. Dentary rectangular and slightly arched with one large and elongate bony
replacement tooth crypt. Symphyseal dentary teeth with well-developed dorsal edge of first
tooth longer than remaining teeth in main row, one circular replacement tooth crypt.
Antorbital narrow, with base wider. Supraorbital narrow, with regular convex ventral margin,
not contacting infraorbital 5. Five well-developed infraorbitals forming semi-circle and
leaving narrow naked area on cheeks. Infraorbitals 3 and 4 with branched sensory canal and 5
with Y-shaped canal. Orbital region broad. Total vertebrae 33 or 34; abdominal vertebrae 14
or 15; caudal vertebrae 19 or 20. One vertebra between verticals through last dorsal-fin
pterygiophore and first anal-fin pterygiophore. Supraneurals 5 or 6.
3.2.6 | Colour in alcohol
Overall body colouration pale to yellowish, dorsal region of body darker. Flanks with
conspicuous rounded or vertically elongate dark blotches. Opercle with a conspicuous dark
blotch on mid region of the bone. Eye with broad dark transverse bar crossing pupil (not
discernible in specimens retained for a long time in formalin). Dorsal, adipose, caudal and
anal fins hyaline, with chromatophores concentrated along distal margin forming narrow

dark stripe. Pectoral and pelvic fins hyaline (Supporting Information Figure S4). Early-stage
juveniles (below 60 mm LS) with mid-lateral ocellus.
3.2.7 | Colour in life
Based on freshly collected specimens. Body overall silver to dark grey, with dorsal region
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darker. Dentary, first two infra-orbitals, inferior portion of head, including preopercle, sub-
opercle, inferior portion of infra-orbitals, opercle and gular, abdominal, supracleithrum and
ventral portion of flank until anal-fin origin orange to light yellow. Opercle with a dark
conspicuous blotch on lower half. Dorsal, adipose, anal and caudal fins hyaline, with dark
chromatophores concentrated on distal portions of fins, forming a dark margin. Pelvic and
anal fin posterior portions red becoming gradually orange to yellow. Pectoral fin in
exemplars larger than 300 mm LS with high concentration of chromatophores (Supporting
Information Figure S5). Juveniles and young adults with strong orange tones on inferior part
of the head, ventral and inferior region of flanks, making them quite similar in colouration to
the red-bellied piranhas (Pygocentrus spp.). The orange intensity decreases in large adults.
3.2.8 | Distribution
Piaractus brachypomus occurs in white-water and clearwater tributaries and occasionally at
the interface of white and black waters. It is widely distributed along the Caquetá-Japurá,
Putumayo-Iça, Branco, Napo, Ucayalí, Solimões, Juruá, Purus, Madeira, Tapajós, Madre de
Dios, Beni, Mamoré, Iténez, Guaporé, Tocantins-Araguaia and Amazonas Rivers in Bolivia,
Brazil, Colombia, Ecuador and Peru (Figure 2).

3.2.9 | Etymology
The name brachypomus refers to the characteristics of the species’ opercle, from the Greek
brachy means width and pomus means cheek.

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4 | DISCUSSION
The present study describes a new species, Piaractus orinoquensis, restricted to the Orinoco
Basin and redescribes the sister species P. brachypomus from the Amazonas River basin,
which was originally described over 200 years ago (Cuvier, 1818). While Machado-Allison
(1982) provided a detailed morphological and ecological analysis of putative P.
brachypomus, differentiating it from Colossoma macropomum, his comparisons were based
on material from the Orinoco River basin and, consequently, in describe P. orinoquensis.
Therefore, our study is the first to provide a detailed description of P. brachypomus, a heavily
exploited commercially important species of the Amazon River basin.
This description was carried out within the integrative taxonomy framework (Padial et
al., 2010) and is based on different but complementary and congruent lines of evidence that
reinforce the conclusions drawn from individual datasets. Escobar et al. (2015) already
demonstrated that Piaractus inhabiting the two basins are distinct biological entities
following independent evolutionary trajectories, having diverged approximately 2.54 M years
ago. Populations from the two basins form reciprocally monophyletic groups and have
distinct body shapes (Figure 2), although these differences are subtle in appearance.
Moreover, these species are diagnosable by the number of scales above and below the lateral
line as well as a suite of molecular character states in the d-loop and coI gene regions (Table

1). All of these lines of evidence support our inference that the Orinoco and Amazon River
basins harbour two different species (Figures 5 and 6).
The third species of the genus, P. mesopotamicus, is restricted to Paraná-Paraguay
River basin (Jégu, 2003; Fricke et al., 2018). Piaractus mesopotamicus possesses
morphological differences that distinguish it from its two congeners, such as tiny scales on
the body, reflected in more lateral line scales (107-128 v. 100 or less), scale rows above and
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below lateral line (49 or more, v. 38 or less); and the lack of a dark opercular blotch in large
individuals (v. presence). However, based on studies by Ortí et al. (2008), Escobar et al.
(2015) and Machado et al. (2018) P. mesopotamicus is phylogenetically nested within P.
brachypomus (Figure 6). In spite of a lack of reciprocal monophyly of P. mesopotamicus and
P. brachypomus, we consider P. mesopotamicus to be a valid species, albeit one that would
benefit from additional taxonomic and phylogenetic analyses.
Our results indicate that the Orinoco and Amazon Basins are occupied by two distinct
species of Piaractus, which raises the question of whether many of the species that are
thought to be shared between the two basins really are conspecific. We predict that analyses
of additional species that are putatively shared between the two basins will reveal many, if
not most, are sister taxa.
4.1 | Management and conservation
Both P. brachypomus and P. orinoquensis sp. nov. are commercially important in the inland
fisheries of the Amazonas and Orinoco Basins (Novoa, 1982; Santos et al., 2006; Agudelo
et al., 2011) (Figure 7). Consequently, it is important to establish independent strategies for
the management of these fisheries and also independent conservation actions that safeguard
the long-term conservation of both species.

As independently evolving lineages, the two species may differ in their life histories,
which would be reflected in population parameters such as birth rate, growth, mortality,
recruitment age, breeding season and population size. These parameters are critical for
establishing management strategies and each species should be regulated by a species-
specific management plan.
Since both species are commonly used in fish-farming and in repopulation and
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reintroduction programmes, these programmes must be developed within the framework of
the existence of two distinct species, each restricted to either the Orinoco Basin or the
Amazon Basin. Otherwise, translocations and release of the non-native species within the
range of the native one can cause hybridisation that, over time, can lead to the extinction of
native species, or simply the displacement of native species and occupation of the new niche
by introduced species (Wolf & Wade, 2001; Allendorf & Luikart, 2007).
This is of particular concern in Colombia, whose national territory encompasses both
the Orinoco and Amazon Basins. The Orinoco River basin is more densely settled and
aquaculture is more intensely developed and based on P. orinoquensis sp. nov. Pisciculture
of P. orinoquensis sp. nov. is practised throughout the country as a food alternative and
source of income for rural low-income communities. Repopulation and reintroduction
programmes are carried out with aquaculture stock, which comprises P. orinoquensis sp.
nov. These repopulation and reintroduction programmes are carried out both in the Orinoco
and the Amazon River basins. However, this type of conservation strategy, which aims to
maintain the long-term viability of fished stocks through increased population size and
through maintaining or increasing genetic diversity, is not fulfilling the proposed goal in
Colombia. Breeding and repopulation programmes in the Putumayo River basin (Colombian
Amazon) are being carried out with P. orinoquensis sp. nov., resulting in P. orinoquensis–P.
brachypomus hybrids being found in nature (Escobar et al., 2015). Interestingly, only F1

hybrids were observed by Escobar et al. (2015), possibly indicating that these hybrids are
sterile.
There is a well-developed theoretical basis for how hybridisation can result in the
extinction of one or both of the species; in the case of repopulation programmes of the rare
local species whose population is being supplemented and numerous empirical examples
showing this (Huff et al., 2011). Even if the extinction of the local species is not a concern
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for management agencies, viable hybrids generally suffer outbreeding depression
(McClelland & Naish, 2006), which also carries an elevated financial cost for management
agencies (Gilk et al., 2004). Thus, well-intentioned programmes may fail, or be
unnecessarily expensive, limiting their effectiveness on the population.
Owing to morphological similarity of the two species, their use in aquaculture and
their susceptibility to translocation and unintentional release, we recommend genetic
monitoring as part of any management strategy, since this is the best tool for delimiting
species and identifying interspecific hybrids. This knowledge, in turn, should be used to
guide management decisions to guarantee the persistence of wild populations of both native
species.
5 | COMPARATIVE MATERIAL EXAMINED
Piaractus mesopotamicus: All from Brazil, Mato Grosso do Sul, Taquarussu. NUP 6638,
119.6 mm LS, lagoa Pousada das Garças, 30 November 2007, Nupélia staff; NUP 9699,
128.3 mm LS, rio Ivinheima, Nupélia Staff, 3 March 2010; NUP 9703, 169.9 mm LS, Canal
Baía, Nupélia staff, 5 March 2010.
ACKNOWLEDGMENTS

We especially thank Adam Leão, Aniello Barbarino, Henry Elias Yucuna Kubeo, Jonathan
Villamil, Luis Francisco Cubillos, Luz Marina Rodriguez, Maria da Conceição Pires, Nirson
Gonzalez and Rosa Elena Martinez Ajiaco for collaborating with field sampling. We are
grateful to Otto Castillo and Oscar León (MCNG), Mariangeles Arce Hernandez and Mark
Sabaj (ANSP) for museum and technical support; to Renata R. Ota (NUP) for analysing
comparative material and to Victória D. P. e Silva (INPA) for preparing the skeletonised
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specimens and X-raying many exemplars. We also thank Otto Enrique Castillo G., Donald C.
Taphorn, Iván Mojica, F. Lima and M. Andrade for their comments and suggestions. Finally,
we are in debt to Mark Sabaj for providing photos in Figures 1, 2, 6, Bruno Barros for photo
in Figure 7, Carlos Lasso and Celsi Señaras for photo in Figure 3, Nadayca Mateussi for
photo Figure 5 and Sergio León for photo in Figure 9b. We are grateful to SISBIO/Brazil,
INCODER/Colombia and INSOPESCA/Venezuela for providing biological sample
collection permits.
CONTRIBUTIONS
M. D. E. contributed to the original idea, development of the study and writing the
manuscript. R. P.O. contributed with morphological analyses and writing the manuscript. A.
M.-A. contributed with morphological data and analyses. J.A. contributed with morphometric
data and analyses. I. P.F. conceived the original idea and contributed with molecular
analyses. T.H. conceived the original idea, supervised the project, analysed data and wrote
the manuscript. All authors discussed the results and contributed to the final manuscript.

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Accepted Article
SUPPORTING INFORMATION
Supporting information can be found in the online version of this paper.

FIGURE 1 Procrustes projections of average body shapes of Piaractus orinoquensis from
the Orininoco Basin (●) and P. brachypomus from the Amzon Basin ( ).
Typesetter
1 Delete and .
2 Change 2x relative to Relative.

Accepted Article
FIGURE 2. Piaractus orinoquensis sp. nov., ANSP 163570, holotype, 150.2 mm standard
length, Laguna La raya, caño Falcón, Portuguesa River, nearby Camaguan, Guárico,
Venezuela.
FIGURE 3. Map of northern South America, showing the distribution of Piaractus
orinoquensis sp. nov. (█; ✩, the type locality; ●, localities of genetic samples; , paratype
localities) and Piaractus brachypomus (█; ●, localities of genetic samples; , localities of
morphologically examined specimens). Symbols may represent more than one lot.
Typesetter
1 Delete .
2 Delete compass arrow.
FIGURE 4 Dry, stuffed and mounted Myletes brachypomus, MNHN A.8627, holotype,
545.0 mm standard length, Brazil.
FIGURE 5 Phylogenetic network of d-loop haplotypes sampled from Piaractus orinoquensis
(●) and P. brachypomus (●).

FIGURE 6 Maximum likelihood phylogenetic tree of the mitochondrial coI barcode region
of representative specimens of Piaractus orinoquensis, P. brachypomus and P.
mesopotamicus. Specimen photos: Piaractus orinoquensis, adult 257 mm standard length
(LS), photo; Piaractus brachypomus, adult 210 LS; Piaractus mesopotamicus, adult 210 LS,;
Mylossoma albiscopum, adult 132 mm LS; Colossoma macropomum, adult 570 LS.
Accepted Article
FIGURE 7 Recently captured adult specimens of: (a) Piaractus orinoquensis, Guaviare
River, tributary of Orinoco River, Colombia; (b) P. orinoquensis, alto Paragua River,
tributary of Caroni River, Venezuela; (c) P. brachypomus upper Solimões River, Puerto
Nariño, Amazonas, Colombia.
Supporting Information
FIGURE S1 Position of landmarks used on geometric morphmetric analyses.
Figure S2 Piaractus orinoquensis sp. nov., ANSP 166685, paratype, 89.8 mm standard
length, Orinoco River, Cabruta, Bolívar, Venezuela. Photo: Mark H. Sabaj.
Figure 3 Piaractus orinoquensis sp. nov., juvenile, 200 mm standard length,, Orinoco River.
Figure S4. Piaractus brachypomus, ANSP 165052, 86.2 mm standard length, Quebrada of
Yanayacu River, W of Miraflores, río Marañon, Loreto, Peru.
Suppl. Figure 5. Piaractus brachypomus adult, UFRO 6289, 315.0 mm standard length, lago
Cuniã, Madeira River, Rondônia, Brazil.

TABLE 1 Analysis of linear distances and meristic counts of Piaractus orinoquensis and P.
brachypomu. Significant differences are highlighted in bold (P < 0.05).
Measurem Mean 95% CI t P Mann- P
_____________________ ___________________ Whitn

ent
____ ey U
____
Accepted Article
P. P. P. P.
orinoquens brachypom orinoquensi brachypomu
is us s s
Proportion
of standard
length
Body 0.590 0.584 0.574–0.606 0.569-0.599 - > 658 >
depth 0.509 0.05 0.05
Predorsal 0.600 0.589 0.594–0.605 0.580-0.598 - > 577 >
length 1.987 0.05 0.05
Prepector 0.328 0.297 0.319–0.337 0.286-0.309 - < 368 <
al length 4.307 0.00 0.00
1 1
Prepelvic 0.549 0.535 0.535–0.564 0.524-0.547 - > 497 <
length 1.577 0.05 0.05
Preanal 0.774 0.757 0.769–0.779 0.750-0.765 - < 417 <
length 3.539 0.00 0.00
1 1
Caudal 0.119 0.113 0.087–0.152 0.110-0.116 - > 1187 <
peduncle 0.376 0.05 0.00
width 1

Caudal 0.116 0.117 0.114–0.118 0.114-0.120 0.677 > 837 >
peduncle 0.05 0.05
depth
Pectoral- 0.234 0.219 0.228–0.234 0.210-0.227 - < 445 <
fin length 2.890 0.01 0.01
Pelvic-fin 0.192 0.173 0.187–0.197 0.166-0.181 - < 346 <
length 4.100 0.00 0.00

Accepted Article
1 1
Dorsal-fin 0.227 0.217 0.216–0.239 0.213-0.221 - > 586 >
base 1.664 0.05 0.05
length
Dorsal-fin 0.251 0.262 0.243–0.260 0.242-0.282 1.023 > 668 >
length 0.05 0.05
Anal-fin 0.283 0.276 0.278–0.288 0.268-0.283 - > 587 >
base 1.659 0.05 0.05
length
Anal-fin 0.205 0.175 0.190–0.220 0.153-0.197 - < 141 <
length 2.391 0.01 0.01
Eye to 0.483 0.481 0.478–0.488 0.475-0.488 - > 695 >
dorsal-fin 0.401 0.05 0.05
origin
Postdorsa 0.539 0.558 0.533–0.545 0.547-0.568 3.117 < 1122 <
l length 0.01 0.00
Dorsal to 0.166 0.153 0.135–0.196 0.147-0.159 - > 777 >
adipose- 0.821 0.05 0.05
fin origin
Adipose- 0.033 0.035 0.031–0.035 0.032-0.037 1.030 > 788 >
fin base 0.05 0.05
length

Head 1.101 1.051 1.077–1.126 0.997-1.105 - < 440 <
membran 1.732 0.01 0.01
ous length
Proportion
off head
length
Accepted Article
Postorbita 0.511 0.505 0.495–0.527 0.491-0.519 -0.589 > 744 >
l length 0.05 0.05
Postorbita 0.605 0.600 0.587–0.623 0.562-0.638 - > 545 >
l 0.242 0.05 0.05
membran
ous
length
Orbital 0.238 0.220 0.228–0.247 0.204-0.237 - > 591 >
diameter 1.851 0.05 0.05
Snout 0.327 0.312 0.317–0.337 0.301-0.323 - 0.04 595 >
length 2.053 4 0.05
Interorbit 0.506 0.487 0.490–0.521 0.477-0.497 - < 479 <
al width 2.046 0.05 0.01
Maxillary 0.215 0.206 0.208–0.221 0197-0.215 - > 660 >
length 0.254 0.251 0.243–0.264 0.243-0.260 1.584 0.05 696 0.05
Eye to - >0.0 >
preopercl 0.385 5 0.05
e distance
Infraorbit 0.155 0.140 0.143–0.166 0.132-0.148 - < 534 <
al length 2.147 0.05 0.05
Head 0.827 0.794 0.801–0.853 0.768-0.820 - > 623 >
depth 1.822 0.05 0.05

Meristics
counts
Dorsal 14.000 15.107 13.879– 14.752- 6.038 < 1249.5 <
rays 14.121 15.462 0.00 0.00
1 1
Pectoral 16.132 16.889 15.922– 16.536- 3.767 < 1037 <

Accepted Article
rays 16.342 17.242 0.00 0.00
1 1
Pelvic 7 7 7–7 7-7 – – 742 NA
rays
Anal rays 24.115 24.357 23.871– 23.849- 0.876 > 761 >
24.360 24.866 0.05 0.05
Lateral 81.906 90.071 80.853– 88.617- 9.260 < 1396 <
line 82.958 91.526 0.00 0.00
scales 1 1
Scales 27.400 34.636 26.806– 32.100- 6.153 < 550 <
above 27.994 37.173 0.00 0.00
lateral 1 1
line
Scales 27.588 34.818 26.917– 33.877- 13.42 < 558 <
below 28.259 35.760 5 0.00 0.00
lateral 1 1
line
Branchial 17.020 16.185 16.791– 15.774- - < 369 <
spines 17.248 16.596 3.627 0.00 0.00
upper 1 1
Branchial 1 1 1–1 1-1 – – 688 NA
spines
lower

Branchial 17.784 17.704 17.432– 17.168- - > 688 >
spines 18.137 18.239 0.257 0.05 0.05
cartilage
Serrae 30.261 31.920 29.205– 30.711- 2.111 > 725 >
prepelvic 31.317 33.129 0.05 0.05
Serrae 20.630 19.840 19.740– 18.145- - > 420 >
postpelvic 21.521 21.536 0.847 0.05 0.05

Accepted Article
Serrae 5.826 5.400 5.465–6.187 4.628-6.172 - > 445 >
bifurcated 1.027 0.05 0.05
Serrae 56.717 57.160 55.155– 54.433- 0.289 > 541 >
total 58.280 59.887 0.05 0.05

TABLE 2 Populations aggregation analysis of genes in Piaractus orinoquensis and P.
brachypomus
Character P. orinoquensis P.brachypomus Character P. orinoquensis P. brachypomus

020 – A 521 T C
077 C – 563 A T
d – Loop
079 T C 633 T C
88 T A 678 T C
101 C T 692 G A
130 A T 736 A –
134 A – 792 – A
Accepted Article
157 G A
163 G A 147 A G
164 A G 150 C T
180 T C 165 C T
188 G A 186 C A
d – Loop
192 T C 222 G A
198 G A 234 C T
212 A G 288 C T
247 T C 334 T C
283 A – coI 339 T C
288 A G 351 C T
453 T – 369 T C
457 T C 435 G A
459 T C 453 C T
468 – T 456 G A
470 T C 522 C T
473 T C 570 A G
478 A G 575 A G
494 T A 576 G G
505 T – 578 – G

TABLE 3 Morphometric data of Piaractus orinoquensis sp. nov.
Measurements n Holotype Paratypes Mean SD
Standard length (mm) 55 150.2 40.0–403.3
Percentages of standard length
Body depth 55 58.6 29.2–67.4 59.3 5.87

Accepted Article
Predorsal length 55 57.9 55.2–65.7 60.1 2.14
Prepectoral length 53 31.4 27.0–41.2 32.2 3.19
Prepelvic length 54 52.3 23.6–60.2 55.7 5.10
Pre-anal length 55 74.5 69.8–81.3 77.3 1.95
Caudal peduncle width 55 10.4 8.7–12.6 10.2 1.00
Caudal peduncle depth 55 11.5 9.8–12.9 11.7 0.75
Pectoral-fin length 54 24.6 18.1–28.6 23.3 1.70
Pelvic-fin length 50 20.0 15.9–23.3 19.0 1.73
Dorsal-fin base length 55 24.1 19.6–24.9 22.3 1.11
Dorsal-fin length 42 30.5 18.6–31.2 25.5 2.82
Anal-fin base length 55 28.6 24.2–31.8 28.5 1.92
Anal-fin length 27 16.5 13.7–28.8 20.2 3.66
Eye to dorsal-fin origin 53 46.2 43.5–53.1 48.3 1.89
Postdorsal length 53 54.0 47.8–61.3 53.9 2.22
Dorsal to adipose-fin origin 54 15.5 11.3–17.9 15.1 1.62
Adipose-fin base length 51 3.6 2.0–4.5 3.3 0.53
Head membranous length 55 33.2 28.9–46.7 36.9 3.98
Head length 55 30.0 20.6–42.5 33.8 3.83
Percentage of head length
Postorbital lenght 55 46.3 39.2–57.5 51.5 3.86

Postorbital membranous length 53 55.3 47.4–67.6 61.1 4.31
Orbital diameter 55 25.2 16.3–29.8 24.1 3.22
Snout length 55 32.5 25.7–38.6 32.5 2.20
Interorbital width 55 53.3 39.0–54.3 50.9 3.43
Maxillary length 55 23.1 15.4–24.3 21.0 1.54
Eye to preopercle distance 53 22.2 18.6–31.1 25.5 2.96

Accepted Article
Infraorbital length 53 18.0 7.0–27.0 15.5 4.04
Head depth 55 38.2 70.9–89.8 81.2 4.46
n, Number of specimens.

TABLE 4 Morphometric data of Piaractus brachypomus.
Measurements n Range Mean SD

Standard length (mm) 28 69.9–675.0
Percentages of standard length

Body depth 28 46.3–64.9 58.6 3.88
Predorsal length 28 52.3–62.5 59.1 2.30
Prepectoral length 28 24.0–35.8 29.8 2.86
Prepelvic length 28 47.0–58.0 53.8 2.88
Pre-anal length 28 71.1–79.1 75.6 2.01
Accepted Article
Caudal peduncle width 28 9.9–13.0 11.2 0.72

Caudal peduncle depth 28 10.1–12.8 11.8 0.64
Pectoral-fin length 28 13.0–25.0 22.0 2.23
Pelvic-fin length 28 13.0–21.8 17.4 1.96
Dorsal-fin base length 28 19.2–23.6 21.8 1.05
Dorsal-fin length 27 17.2–38.1 27.4 4.97
Anal-fin base length 28 23.7–31.9 27.5 1.85
Anal-fin length 19 10.1–26.5 16.1 4.42
Eye to dorsal-fin origin 28 44.8–51.1 48.2 1.60
Postdorsal length 28 48.3–63.6 55.9 2.71
Dorsal to adipose-fin origin 28 13.0–17.7 15.2 1.44
Adipose-fin base length 28 2.3–5.4 3.4 0.72
Head membranous length 28 26.5–38.6 34.1 3.14
Head length 28 25.2–35.7 31.6 2.66
Percentages of head length
Postorbital length 28 44.5–56.8 51.4 2.90
Postorbital membranous length 28 51.8–65.8 57.9 3.43
Orbital diameter 28 13.0–29.5 23.4 4.04
Snout length 28 24.6–35.5 32.1 2.81
Interorbital width 28 43.6–53.1 48.9 2.53
Maxillary length 28 16.0–24.2 20.7 2.27
Eye to preopercle distance 28 18.6–29.5 25.5 2.09
Infraorbital length 28 9.6–18.1 13.7 1.99
Head depth 28 70.5–87.0 78.6 5.50
n, Number of specimens.

Figure 1
Accepted Article
Figure
Piaractus shape
Orinoco
Amazon
6e+07
2e+07
-2e+07
relative body depth
Accepted Article
-6e+07
-1e+08 -5e+07 0e+00 5e+07 1e+08
relative body length
Figure
Accepted Article This article is protected by copyright. All rights reserved.
Figure
Accepted Article
Figure
5 ind
Orinoco Amazon
30 bp
Accepted Article
Figure
Mylossoma albiscopum
Amazon River basin Piaractus brachypomus GuajaraMirim2
Piaractus brachypomus LaPedrera1 Piaractus brachypomus
Piaractus mesopotamicus LBP23804 Amazon River basin
Piaractus brachypomus Solimoes1
Piaractus brachypomus Solimoes2
Piaractus brachypomus Santarem10
Piaractus brachypomus Santarem11
Piaractus mesopotamicus CEPTA003
Piaractus brachypomus Janauaca1
Piaractus brachypomus Branco3
Accepted Article

Piaractus brachypomus GuajaraMirim1
Piaractus brachypomus Itaituba2
Colossoma macropomum Piaractus mesopotamicus CEPTA005
Amazon River basin Piaractus brachypomus N192 Piaractus mesopotamicus
Piaractus brachypomus Itaituba1
Piaractus brachypomus LaPedrera2 Parana/Paraguay River basin
Piaractus mesopotamicus LBP23805
100 Piaractus brachypomus CEPTA007
Piaractus brachypomus CEPTA014
Piaractus brachypomus Branco5
Piaractus brachypomus Janauaca2
Piaractus mesopotamicus OM20
Piaractus orinoquensis Meta11
Piaractus orinoquensis Guaviare16
100 Piaractus orinoquensis Meta8 Piaractus orinoquensis
Orinoco River basin
Piaractus orinoquensis Apure1
Piaractus orinoquensis Guaviare15
Piaractus orinoquensis SanFelixV1
Piaractus orinoquensis Apure3
100 Piaractus orinoquensis Meta9
Piaractus orinoquensis SanFelix3
Colossoma macropomum
Mylossoma albiscopum
0.02
Figure
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A new species of Piaractus (Characiformes: Serrasalmidae) from the Orinoco

Article in Journal of Fish Biology · April 2019

Maria Doris Escobar Lizarazo Rafaela Ota

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Izeni Pires Farias Tomas Hrbek

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with a redescription of Piaractus brachypomus

M. D. ESCOBAR1,2 | R. P. OTA3 | A. MACHADO-ALLISON4,5 | | I. P. FARIAS1 | T.

Coroado I., Manaus, Amazonas, Brasil.

Sistemática de Peces, Ciudad Universitaria de Caracas, Los Chaguaramos, Caracas,

M. D. Escobar, Universidade Federal do Amazonas, Departamento de Genética, Instituto de

Ciências Biológicas, Laboratório de Evolução e Genética Animal, Av. General Rodrigo

Octávio, 6200, Coroado I. Cep: 69080-90, Manaus, Amazonas, Brasil.

This article is protected by copyright. All rights reserved.

Cientifico e Tecnológico (CNPq 490682/2010–3 and 482662/2013–1 to T.H.; CNPq/PPG7

5570090/2005–9 and CNPq/CT-Amazon 575603/2008–9 to I.P.F.), Coordenação de

Aperfeiçoamento de Pessoal de Nível Superior (CAPES 53923790287) PhD fellowship to

(306804/2013-3) and T.H. (303646/2010-1).

provide a re-description of Piaractus brachypomus, restricting its geographic distribution to

and need to be independently managed as distinct species.

Amazon Basin, meristics, mitochondrial DNA, morphology, serrasalmid

This article is protected by copyright. All rights reserved.

(Berg 1895), within his treatment of P. brachypomus. Subsequently, Machado-Allison

species in the Madeira River (Brazilian Amazon).

Géry (1985) highlighted the extensive range of intraspecific phenotypic variability,

brachypomus and P. mesopotamicus. However, in spite of Géry’ s (1985) observations, all

and distribution of P. brachypomus proposed by Machado-Allison (1982). Escobar et al.

This article is protected by copyright. All rights reserved.

Traditionally, species descriptions are based primarily on morphological characters,

Given the increasingly easy accessibility of molecular techniques to taxonomists, an

2005) can provide robust taxonomic hypotheses, especially in morphologically conserved

reciprocal monophyly, mutually exclusive mating systems and diagnosability. In addition to

This article is protected by copyright. All rights reserved.

of P. brachypomus and we delimit the geographic distribution of both species.

2 | MATERIAL AND METHODS

2.1 | Fish samples

the ichthyology reference collection of the Guyana Hydrobiological Research Station, La

Central University of Venezuela (MBUCV-V), Caracas, Venezuela.

(QGIS Development Team, 2018).

2.2 | Molecular methods

This article is protected by copyright. All rights reserved.

from the Amazon Basin (GenBank KP705146–KP705254) and 33 cytochrome c oxidase

from the Amazon Basin (Genbank KP723341–KP723360, MK395376– MK395388).

software Haploviewer (Salzburger et al., 2011) and a maximum-likelihood phylogeny

We also inferred a maximum-likelihood phylogeny from the coI sequence alignment in

outgroups we used Colossoma macropomum (Cuvier 1816) (MG751942) and Mylossoma

albiscopum (Cope 1872) (MG752455) and we included P. mesopotamicus as additional

coalescent single-locus species-discovery method that has been designed specifically to

reduced the dataset to unique haplotypes using the function hapCollapse

(www.github.com/legalLab/protocols-scripts) to further minimise interpreting population-

This article is protected by copyright. All rights reserved.

2.3 | Morphological methods

standard methods proposed in Machado-Allison (1982). All distances were calculated in mm

and transformed to percentages; the body measurements are expressed as a proportion of

basins. These analyses were performed in the STATS package in R.

Vertebral and supraneural elements were counted in 17 skeletonised specimens [eight

of P. brachypomus: INPA 46334 (1 skeleton), 46335 (1 skeleton), 50841 (4 skeletons), 57799

This article is protected by copyright. All rights reserved.

and Mattox et al. (2014).

In the descriptions, counts are followed by their frequency in parentheses and an

abbreviations are: IAvH, Instituto de Investigación de Recursos Biológicos Alexander von

Humboldt, Villa de Leyva, Boyacá, Colombia; ICN-MHN, Instituto de Ciencias Naturales,